Selective cytotoxicity and modification activity of picornaviruses on transformed cell lines

Aim. We do analyze of the dynamics of morphometabolic changes in transformed cells (of susceptoible lines) demonstrating resistance to picrnaviral infection. Methods. The study was performed by application of cell culture technology and a complex of cytochemical and cytophotometric assays. Were us...

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Datum:	2011
Hauptverfasser:	Karalyan, Z.A., Avagyan, H.R.
Format:	Artikel
Sprache:	English
Veröffentlicht:	Інститут молекулярної біології і генетики НАН України 2011
Schriftenreihe:	Вiopolymers and Cell
Online Zugang:	http://dspace.nbuv.gov.ua/handle/123456789/156377
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spelling	irk-123456789-1563772019-06-19T01:25:18Z Selective cytotoxicity and modification activity of picornaviruses on transformed cell lines Karalyan, Z.A. Avagyan, H.R. Aim. We do analyze of the dynamics of morphometabolic changes in transformed cells (of susceptoible lines) demonstrating resistance to picrnaviral infection. Methods. The study was performed by application of cell culture technology and a complex of cytochemical and cytophotometric assays. Were used picornaviruses from various genu. Results. According to the results obtained, resistant to picornavirus infection cells of different susceptible lines have similar changes in the phenotype. They have decreased number of nucleoli and increased percentage of euploidy (and near euploid). In resistant cells of all cultures the reduction in amount of DNA and RNA both in nucleus and in cytoplasm wasfound. All these data correlated with the increased euploidy (and near euploid) of the resistant population. All picornavirus resistant cells had a less transformed phenotype, and decreased proliferative activity. Decreased nucleolar status becomes apparent by reduction of all nucleolar indices. Conclusions. Picornaviruses on the susceptible cells produce 2 types of changes – selection and modification. Whateverthe mechanism, it isspecific for an individual virus, since no restrictions occur in case of infection caused by another picornavirus. Keywords: picornavirus, euploidy, nucleus, nucleolus. Мета. Мета даної роботи полягала у вивченні динаміки морфологічних і фізіологічних змін трансформованих клітин, резистентних до пікорнавірусної інфекції. Методи. Дослідження виконано за умов in vitro із застосуванням цитохімічного і цитофотометричного аналізу. У роботі використано пікорнавіруси різних родів. Результати. Встановлено, що стійкі до інфікування пікорнавірусами клітини різних чутливих ліній набувають аналогічних змін у фенотипі. У резистентних клітинах усіх культур як у ядрі, так і цитоплазмі виявлено зниження вмісту ДНК і РНК. Усі ці дані корелюють з підвищенням еуплоїдної (та біляеуплоїдної) популяції за формування резистентності. Усі резистентні до пікорнавірусів клітини були меншими за розмірами порівняно з початковим трансформованим фенотипом та демонстрували зниження проліферативної активності. Зменшення активності ядерець супроводжується вірогідним падінням усіх ядерцевих показників. Висновки. Пікорнавіруси проявляють подвійну дію на чутливі клітини, яка виражається у селективній цитотоксичності і модифікуючому впливі. При цьому механізми їхньої дії є специфічними для кожного окремого пікорнавірусу. Так, резистентні стосовно одного пікорнавірусу клітини виявляються нестійкими до інфекції, спричиненої іншими пікорнавірусами. Ключові слова: пікорнавіруси, еуплоїдія, ядро, ядерце. Цель. Целью данной работы явилось изучение динамики морфо - логических и физиологических изменений трансформированных клеток, резистентных к пикорнавирусной инфекции. Методы. Исследование проведено в условиях in vitro с применением цитохимического и цитофотометрического анализов. В работе использованы пикорнавирусы различных родов. Результаты. Установлено, что устойчивые к инфицированию пикорнавирусами клетки различных чувствительных линий приобретают аналогичные изменения в фенотипе. В резистентных клетках всех культур как в ядре, так и цитоплазме выявлено снижение содержания ДНК и РНК. Все эти данные коррелируют с повышением эуплоидной (и околоэуплоидной) популяции при формировании резистентности. Все резистентные к пикорнавирусам клетки были меньше по размерам в сравнении с начальным трансформированным фенотипом и демонстрировали снижение пролиферативной активности. Уменьшение ядрышковой активности сопровождается достоверным падением всех ядрышковых показателей. Выводы. Пикорнавирусы проявляют двойное действие на чувствительные клетки, выражающееся в селективной цитотоксичности и модифицирующем влиянии. При этом механизмы их действия являются специфичными для каждого отдельного пикорнавируса. Так, резистентные в отношении одного пикорнавируса клетки оказываются неустойчивыми к инфекции, вызванной другими пикорнавирусами. Ключевые слова: пикорнавирусы, эуплоидия, ядро, ядрышко. 2011 Article Selective cytotoxicity and modification activity of picornaviruses on transformed cell lines / Z.A. Karalyan, H.R. Avagyan // Вiopolymers and Cell. — 2011. — Т. 27, № 5. — С. 377-380. — Бібліогр.: 8 назв. — англ. 0233-7657 DOI: http://dx.doi.org/10.7124/bc.000127 http://dspace.nbuv.gov.ua/handle/123456789/156377 578.821.5:615.371 en Вiopolymers and Cell Інститут молекулярної біології і генетики НАН України
institution	Digital Library of Periodicals of National Academy of Sciences of Ukraine
collection	DSpace DC
language	English
description	Aim. We do analyze of the dynamics of morphometabolic changes in transformed cells (of susceptoible lines) demonstrating resistance to picrnaviral infection. Methods. The study was performed by application of cell culture technology and a complex of cytochemical and cytophotometric assays. Were used picornaviruses from various genu. Results. According to the results obtained, resistant to picornavirus infection cells of different susceptible lines have similar changes in the phenotype. They have decreased number of nucleoli and increased percentage of euploidy (and near euploid). In resistant cells of all cultures the reduction in amount of DNA and RNA both in nucleus and in cytoplasm wasfound. All these data correlated with the increased euploidy (and near euploid) of the resistant population. All picornavirus resistant cells had a less transformed phenotype, and decreased proliferative activity. Decreased nucleolar status becomes apparent by reduction of all nucleolar indices. Conclusions. Picornaviruses on the susceptible cells produce 2 types of changes – selection and modification. Whateverthe mechanism, it isspecific for an individual virus, since no restrictions occur in case of infection caused by another picornavirus. Keywords: picornavirus, euploidy, nucleus, nucleolus.
format	Article
author	Karalyan, Z.A. Avagyan, H.R.
spellingShingle	Karalyan, Z.A. Avagyan, H.R. Selective cytotoxicity and modification activity of picornaviruses on transformed cell lines Вiopolymers and Cell
author_facet	Karalyan, Z.A. Avagyan, H.R.
author_sort	Karalyan, Z.A.
title	Selective cytotoxicity and modification activity of picornaviruses on transformed cell lines
title_short	Selective cytotoxicity and modification activity of picornaviruses on transformed cell lines
title_full	Selective cytotoxicity and modification activity of picornaviruses on transformed cell lines
title_fullStr	Selective cytotoxicity and modification activity of picornaviruses on transformed cell lines
title_full_unstemmed	Selective cytotoxicity and modification activity of picornaviruses on transformed cell lines
title_sort	selective cytotoxicity and modification activity of picornaviruses on transformed cell lines
publisher	Інститут молекулярної біології і генетики НАН України
publishDate	2011
url	http://dspace.nbuv.gov.ua/handle/123456789/156377
citation_txt	Selective cytotoxicity and modification activity of picornaviruses on transformed cell lines / Z.A. Karalyan, H.R. Avagyan // Вiopolymers and Cell. — 2011. — Т. 27, № 5. — С. 377-380. — Бібліогр.: 8 назв. — англ.
series	Вiopolymers and Cell
work_keys_str_mv	AT karalyanza selectivecytotoxicityandmodificationactivityofpicornavirusesontransformedcelllines AT avagyanhr selectivecytotoxicityandmodificationactivityofpicornavirusesontransformedcelllines
first_indexed	2025-07-14T06:25:15Z
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fulltext	Selective cytotoxicity and modification activity of picornaviruses on transformed cell lines Z. A. Karalyan, H. R. Avagyan Institute of Molecular Biology, NAS of Republic of Armenia 7, Hasratyan Str., Yerevan, Republic of Armenia, 0014 zkaralyan@yahoo.com Aim. We do analyze of the dynamics of morphometabolic changes in transformed cells (of susceptoible lines) demonstrating resistance to picrnaviral infection. Methods. The study was performed by application of cell culture technology and a complex of cytochemical and cytophotometric assays. Were used picornaviruses from various genu. Results. According to the results obtained, resistant to picornavirus infection cells of different susceptible lines have similar changes in the phenotype. They have decreased number of nucleoli and increased percentage of euploidy (and near euploid). In resistant cells of all cultures the reduction in amount of DNA and RNA both in nucleus and in cytoplasm was found. All these data correlated with the increased euploidy (and near euploid) of the resistant population. All picornavirus resistant cells had a less transformed phenotype, and decreased proliferative activity. Decreased nucleolar status becomes apparent by reduction of all nucleolar indices. Conclusions. Picornaviruses on the susceptible cells produce 2 types of changes – selection and modification. Whatever the mechanism, it is specific for an individual virus, since no restrictions occur in case of infection caused by another picornavirus. Keywords: picornavirus, euploidy, nucleus, nucleolus. Introduction. Although selection for host cells that resist viral infection during picornaviral persistence has been reported, their role in persistence has not been elucidated, and the basis for host cell resistance is not clear [1, 2]. We analyze the possibility of complex me- tabolism changes in resistant cells as a protecting factor against virus infection. Aim of this research is the analysis of the dynamics of morphometabolic peculiarities of the becoming re- sistant cells as well as the comparison of phenotype of picornavirus resistant cell of sensitive to the cytotoxic activity lines. Materials and methods. Cells. HEp-2, HEK293, Caco-2, BHK cells, continuous transformed cultures, were cultivated as described (ATCC). Viruses. EMCV (Columbia-SK strain) was used at multiplicity of infection 0.1 TCD50 per cell. Poliovirus- 1 /Sabin/: was used at multiplicity of infection 0.1 TCD50 per cell. Foot-and-mouse disease virus (strains O NKR-194; A-22; Asia-1 – FMDV-O, FMDV-A, FMDV-Asia) was used at multiplicity of infection 0.1 TCD50 per cell. Viral titers were calculated by the me- thod of Karber. As a control the parallel conducted pas- sages of noninfected cultures were used. After lytic viral infection in some flaks were pre- sent isolated cells which have resistance against corres- ponding virus. Resistant cells were received by one ti- me infection. The second method of receiving the resistant cells was described by Taber et al. [3]. Chronic viral infec- tion was received by one time infection of 48-hour mo- nolayers of cultures CaCo-2 and НЕр-2 by poliovirus-1 /Sabin/. The multiplicity of infection was 0.000001 TCD50 per cell. Infected cells were incubated at 36.5– 37 oC. After 5–11 passages some cultures become free from virus and cells received resistance to a correspon- ding virus. Chronic infections were repeated 3 times and summarized data were presented. 377 ISSN 0233–7657. Biopolymers and Cell. 2011. Vol. 27. N 5. P. 377–380  Institute of Molecular Biology and Genetics, NAS of Ukraine, 2011 Cytochemical and cytophotometric analysis. For si- multaneously staining DNA and RNA the methyl green, pyronin Y method was used. DNA quantification was do- ne by Feulgen staining. For quantification of RNA was used gallocyanin chromalum stain. In gallocyanin chroma- lum preparation the control data were defined as 100 %. Cell viability test. Was used the trypan blue exclu- sion test, cells were incubated for 5 min with 0.04 % trypan blue («Sigma», USA). Results and discussion. The received chronically infected cultures were characterized by insular and slo- wer growth (approximately twice), lack of the capa- city to form monolayer and constant decreased produc- tion of a virus. After 5–11 passages some cultures beca- me free from poliovirus and cells received resistance to it. In microscopic examination of chronically infected cultures we revealed a gradual change towards small round cell morphology. The received resistant cells survived under the in- fection with analogical virus, but did not survive under the infection with other viruses. Proliferative activity of the resistant to picornavirus cells of susceptible cultures was lower than that of the cells of control populations. It happens due to about 2– 3-fold increase of the interphase of mitotic cycle com- pared to the control populations. According to the results of cytophotometric analy- sis of the acute picornaviral infection of Hela, CaCo-2, HEK293, RD and HEp-2 cultures had similar morpho- logical changes. The survived cells had similar pheno- typical characteristics. This was evidenced by the pre- sence of binucleate cells, and the decreased number of nucleoli in each nucleus, which is visible using MGP staining. Those cells were smaller and had a round sha- pe, in comparison with the control ones. The reduction in sizes applies to both the cells and theirs nuclei. The resistant cells had significantly decreased DNA amo- unt (Table 1). The similar changes were noticed under the influence of chronical picornaviral infection. The nucleoli number in investigated cells was redu- ced compared to the control population, which testifies the oppression of translational activity (Table 2). The percent of euploid cells among the resistant cells in RD, Hep-2, HEK293, BHK-21 and HeLa, signifi- cantly increased under the acute infection of EMCV. In case of an acute and chronic poliovirus infection among the resistant cells in Hep-2 cells the percentage of eu- ploid cells also significantly increased, whereas in the CaCo-2 cells such changes are insignificant (Table 3). The disparity in viral effects observed in CaCo-2 cells can be explained by diverse levels of differentiation [4]. It can be noticed that in all cases of resistant cells, the number of nucleoli in the nuclei has tendency to de- crease. This data correlates with the increased per- centage of euploid cells in resistant populations. The reduction in the nucleoli number testifies oppression of metabolic activity in common and translational activity in particular. The structure of euploid population also changed. In control population of CaCo-2, RD and Hep-2 cells prevailed over the tetraploid cells with a few number of octaploid and 16 «c» cells. In resistant to picornavirus cells appeared a great deal of diploid cells and disappea- red the population of 8 «c» and 16 «c» cells. In resistant HEK293 cells the tetraploid population increased, the oc- tapoid disappeared and in HeLa aroused diploid cells. In resistant cells of all cultures is found the reducti- on of DNA, RNA amount, in both nucleus and cyto- 378 KARALYAN Z. A., AVAGYAN H. R. Cultures Control «c» EMCV acute infection «с» Poliovirus acute infection «с» Poliovirus chronic infection «с» FMDV-O acute infection «с» FMDV-A acute infection «с» FMDV-Asia acute infection «с» RD 3.9 ± 0.6 2.5 ± 0.2* – – – – – Hep-2 4.4 ± 0.4 3.4 ± 0.2* 3.8 ± 0.2* 3.6 ± 0.1* – – – HEK293 5.9 ± 0.8 4.1 ± 0.5 – – – – – HeLa 5.7 ± 0.8 4.6 ± 0.8 4.0 ± 0.4 – – – – CaCo-2 5.6 ± 1.0 – 3.2 ± 0.7 4.1 ± 0.4 – – – BHK-21 2.8 ± 0.4 – – – 2.0 ± 0.8 3.0 ± 0.2 1.8 ± 0.1* Significant compared to control, p < 0.05–p < 0.01; t = 1.88; t = 1.89. Table 1 Amount of DNA in control, resistant cells of the RD, Hep-2, HEK293, HeLa, CaCo-2, BHK-21 cultures («c» units) plasm. All these data correlated with the increased eu- ploidy of the resistant population. In the meantime, the percentage of euploidy in a cell culture at control and vi- ral infection is closely invert correlated to the number of nucleoli. So resistant cells of all cell lines such as HEK293, RD, HeLa, HEp-2, showed decreased num- ber of nucleoli and increased percentage of euploidy. The percentage of euploid cells resistant to the picorna- viral infection increases in all cultures (except CaCo-2). We can also conclude that picornaviruses not only do selective action but also can modify the cells. Modi- fication consisted of the occurrence in survived cells (RD, Hep-2) of diploid population absent in control. In favour of modification testify the changes of cell pheno- type that have survived after an acute and chronic infec- tion of picornaviruses. The main selective factor is apoptosis induced in the infected cells. Modification of cells occurs by deblocking the cells in phase G2 and by stimulation of their division. In general, the phenotype of resistant cells can be characterized as less transfor- med compared with the intact cell populations. Resuming the action of picornaviruses on the sus- ceptible cells we can define 2 types of changes – selec- tion and modification. Possibly, the viruses induced apoptosis selectively in multinucleolar aneuploid cells. In favour of this assumption testify the data obtained by Taylor, Martin-DeLeon [5] that the number of nucleo- lar-forming regions is genotypically determined, so dif- ferences among subgroups of multinucleolar cells are more likely than the production of new clones. By La- badie et al. [6] was shown that CaCo-2 cells which are partially resistant to poliovirus induced apoptosis can be selected during persistent virus infection. Castedo et al. [7] described that diploid cells were more stable to the apoptotic influence. Although the presence or absence of virus receptors on the cell surface remains a major determining factor of the susceptibility of a cell to virus infection, there is now incre- asing evidence that the intracellular environment plays an important role in the outcome of viral invasion [8]. Important metabolic characteristic of resistant to pi- cornavirus infection cells is a significant decrease in cel- 379 SELECTIVE CYTOTOXICITY AND MODIFICATION ACTIVITY OF PICORNAVIRUSES Cultures Control EMCV acute infection Poliovirus acute infection Poliovirus chronic infection FMDV-O acute infection FMDV-A acute infection FMDV-Asia acute infection RD 2.7 ± 0.3 1.6 ± 0.1 – – – – – Hep-2 2.6 ± 0.2 1.3 ± 0.2* 1.6 ± 0.1* 1.5 ± 0.1* – – – HEK293 2.3 ± 0.2 2.0 ± 0.1 – – – – – HeLa 2.5 ± 0.2 1.9 ± 0.2* 2.0 ± 0.2 – – – – CaCo-2 2.9 ± 0.3 – 2.1 ± 0.1* 2.0 ± 0.2* – – – BHK-21 2.1 ± 0.2 – – – 1.1 ± 0.1* 1.1 ± 0.1* 0.9 ± 0.1* Significant compared to control, p < 0.05–p < 0.01. Table 2 Changes in nucleoli number in control and resistant cells of the RD, Hep-2, HEK293, HeLa, CaCo-2 cultures Cultures Control, % EMCV acute infection, % Poliovirus acute infection, % Poliovirus chronic infection, % FMDV-O acute infection FMDV-A acute infection, % FMDV-Asia acute infection, % RD 20.4 ± 3.5 35.4 ± 5.9 – – – – – Hep-2 13.0 ± 4.1 24.2 ± 3.8* 31.0 ± 7.7* 34.0 ± 5.1* – – – HEK293 26.8 ± 3.8 42.1 ± 4.3* – – – – – HeLa 23.6 ± 3.2 35.5 ± 5.8 32.8 ± 3.3* – – – – CaCo-2 50.1 ± 4.1 – 48.2 ± 6.8 57.5 ± 7.3 – – – BHK-21 22.0 ± 5.2 – – – 33.0 ± 4.9 46.2 ± 8.7 42.8 ± 6.0 *Significant compared to control, p < 0.05–p < 0.01. Table 3 Euploidy in control and resistant cells of the RD, Hep-2, HEK293, HeLa, CaCo-2, BHK-21 cultures (%) lular RNA amount. This phenomenon is described in all investigated parts of cells – nucleoli, nucleus, cyto- plasm. Undoubtedly first of all the synthesis of rRNA, the major part of total cellular RNA, is oppressed. All picornavirus resistant cells had a less transformed phe- notype, such as decreased proliferative activity, redu- ced DNA amount, increased euploid population and de- creased nucleolar status. Decreased nucleolar status be- comes apparent by reduction of absolute and relative nucleolar indices. Consequently the viral titers reduc- tion in resistant cells could be the direct result of dimi- nished activity of the RNA synthesis machinery. Whatever the mechanism, it is specific for individual virus since no restrictions occur in infections with other picornaviruses. З. А. Ка ра лян, Г. Р. Авагян Се лек тив на ци то ток сичність і мо дифіку ю ча дія пікор навірусів на транс фор мо ва них клітин них лініях Ре зю ме Мета. Мета да ної ро бо ти по ля га ла у вив ченні ди наміки мор фо ло- гічних і фізіологічних змін транс фор мо ва них клітин, ре зис тент- них до пікор навірус ної інфекції. Ме то ди. Досліджен ня ви ко на но за умов in vitro із за сто су ван ням ци тохімічно го і ци то фо то мет - рич но го аналізу. У ро боті ви ко рис та но пікор навіруси різних ро- дів. Ре зуль та ти. Вста нов ле но, що стійкі до інфіку ван ня пікор - навіру са ми клітини різних чут ли вих ліній на бу ва ють ана логічних змін у фе но типі. У ре зис тен тних кліти нах усіх куль тур як у ядрі, так і ци топ лазмі ви яв ле но зни жен ня вмісту ДНК і РНК. Усі ці дані ко ре лю ють з підви щен ням еуп лої дної (та біля е уп лої дної) по - пу ляції за фор му ван ня ре зис тен тності. Усі ре зис тентні до пі- кор навірусів клітини були мен ши ми за розмірами порівня но з по - чат ко вим транс фор мо ва ним фе но ти пом та де мо нстру ва ли зни - жен ня проліфе ра тив ної ак тив ності. Змен шен ня ак тив ності яде- рець суп ро вод жується вірогідним падінням усіх ядер це вих по каз - ників. Вис нов ки. Пікор навіруси про яв ля ють подвійну дію на чут - ливі клітини, яка ви ра жається у се лек тивній ци то ток сич ності і мо дифіку ю чо му впливі. При цьо му ме ханізми їхньої дії є спе ци- фічни ми для кож но го окре мо го пікор навірусу. Так, ре зис тентні сто сов но од но го пікор навірусу клітини ви яв ля ють ся нестійки ми до інфекції, спри чи не ної інши ми пікор навіру са ми. Клю чові сло ва: пікор навіруси, еуп лоїдія, ядро, ядер це. З. А. Ка ра лян, Г. Р. Авагян Се лек тив ная ци то ток сич ность и мо ди фи ци ру ю щее де йствие пи кор на ви ру сов на транс фор ми ро ван ных кле точ ных ли ни ях Ре зю ме Цель. Целью дан ной ра бо ты яви лось из уче ние ди на ми ки мор фо - ло ги чес ких и фи зи о ло ги чес ких из ме не ний транс фор ми ро ван ных кле ток, ре зис тен тных к пи кор на ви рус ной ин фек ции. Ме то ды. Иссле до ва ние про ве де но в усло ви ях in vitro с при ме не ни ем ци то - хи ми чес ко го и ци то фо то мет ри чес ко го ана ли зов. В ра бо те ис - поль зо ва ны пи кор на ви ру сы раз лич ных ро дов. Ре зуль та ты. Ус- та нов ле но, что устой чи вые к ин фи ци ро ва нию пи кор на ви ру са ми клет ки раз лич ных чу встви тель ных ли ний при об ре та ют ана ло - гич ные из ме не ния в фе но ти пе. В ре зис тен тных клет ках всех куль тур как в ядре, так и ци топ лаз ме вы яв ле но сни же ние со дер - жа ния ДНК и РНК. Все эти дан ные кор ре ли ру ют с по вы ше ни ем эуп ло ид ной (и око ло э уп ло ид ной) по пу ля ции при фор ми ро ва нии ре - зис тен тнос ти. Все ре зис тен тные к пи кор на ви ру сам клет ки были мень ше по раз ме рам в срав не нии с на чаль ным транс фор ми - ро ван ным фе но ти пом и де мо нстри ро ва ли сни же ние про ли фе ра - тив ной ак тив нос ти. Умень ше ние яд рыш ко вой ак тив нос ти со про вож да ет ся дос то вер ным па де ни ем всех яд рыш ко вых по ка - за те лей. Вы во ды. Пи кор на ви ру сы про яв ля ют двой ное де йствие на чу встви тель ные клет ки, вы ра жа ю ще е ся в се лек тив ной ци то - ток сич нос ти и мо ди фи ци ру ю щем вли я нии. При этом ме ха низ мы их де йствия яв ля ют ся спе ци фич ны ми для каж до го от дель но го пи кор на ви ру са. Так, ре зис тен тные в от но ше нии од но го пи кор на - ви ру са клет ки ока зы ва ют ся не устой чи вы ми к ин фек ции, вы зван - ной дру ги ми пи кор на ви ру са ми. Клю че вые сло ва: пи кор на ви ру сы, эуп ло и дия, ядро, яд рыш ко. REFERENCES 1. Borzakian S., Couderc T., Barbier Y., Attal G., Pelletier I., Col- bere-Garapin F. Persistent poliovirus infection: establishment and maintenance involve distinct mechanisms // Virology.– 1992.–186, N 2.–P. 398–408. 2. Martin Hernandez A. M., Carrillo E. C., Sevilla N., Domingo E. Rapid cell variation can determine the establishment of a persis- tent viral infection // Proc. Natl Acad. Sci. USA.–1994.–91, N 9.–P. 3705–3709. 3. Taber R., Alexander V., Whitford W. Persistent reovirus infecti- on of CHO cells resulting in virus resistance // J. Virol.–1976.– 17, N 2.–P. 513–524. 4. Karalyan Z. A., Jaghatspanyan N. G., Gasparyan M. H., Ha- kobyan L. A., Abroyan L. O., Ter-Pogossyan Z. R., Kamalyan L. A., Karalova E. M. Comparison of impact of EMCV replication on the nuclear apparatus of NIH 3T3 and HEp-2 cells // Cell Biol. International.–2005.–29, N 7.–P. 586–592. 5. Taylor E. F., Martin-DeLeon P. A. Familial silver staining pat- terns of human nucleolus organizer regions (NORs) // Am. J. Hum. Genet.–1981.–33, N 1.–P. 67–76. 6. Labadie K., Saulnier A., Martin-Latil S., Colbere-Garapin F. Reduced apoptosis in human intestinal cells cured of persistent poliovirus infection // J. Virol.–2007.–81, N 6.–P. 3033–3036. 7. Castedo M., Coquelle A., Vivet S., Vitale I., Kauffmann A., Des- sen P., Pequignot M. O., Casares N., Valent A., Mouhamad S., Schmitt E., Modjtahedi N., Vainchenker W., Zitvogel L., Lazar V., Garrido C., Kroemer G. Apoptosis regulation in tetraploid cancer cells // EMBO J.–2006.–25, N 11.–P. 2584–2595. 8. Strong J. E., Coffey M. C., Tang D., Sabinin P., Lee P. W. The molecular basis of viral oncolysis: usurpation of the Ras sig- naling pathway by reovirus // EMBO J.–1998.–17, N 12.– P. 3351–3362. UDC 578.821.5:615.371 Received 27.07.11 380 KARALYAN Z. A., AVAGYAN H. R.

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